A Synopsis of the Genera and Species in the Tylenchorhynchinae (Tylenchoidea, Nematoda)1

OF WASHINGTON, VOLUME 40, NUMBER 1, JANUARY 1973 123

Speer, C. A., and D. M. Hammond. 1970. tured bovine cells. J. Protozool. 18 (Suppl.): Development of Eimeria larimerensis from the 11. Uinta ground squirrel in cell cultures. Ztschr. Vetterling, J. M., P. A. Madden, and N. S. Parasitenk. 35: 105-118. Dittemore. 1971. Scanning electron mi- , L. R. Davis, and D. M. Hammond. croscopy of poultry coccidia after in vitro 1971. Cinemicrographic observations on the excystation and penetration of cultured cells. development of Eimeria larimerensis in cul- Ztschr. Parasitenk. 37: 136-147.

A. C. TARJAN2

ABSTRACT: The genera Uliginotylenchus Siddiqi, 1971, Quinisulcius Siddiqi, 1971, Merlinius Siddiqi, 1970, Ttjlenchorhynchus Cobb, 1913, Tetylenchus Filipjev, 1936, Nagelus Thome and Malek, 1968, and Geocenamus Thorne and Malek, 1968 are discussed. Keys and diagnostic data are presented. The following new combinations are made: Tetylenchus aduncus (de Guiran, 1967), Merlinius alboranensis (Tobar-Jimenez, 1970), Geocenamus arcticus (Mulvey, 1969), Merlinius brachycephalus (Litvinova, 1946), Merlinius gaudialis (Izatullaeva, 1967), Geocenamus longus (Wu, 1969), Merlinius parobscurus ( Mulvey, 1969), Merlinius polonicus (Szczygiel, 1970), Merlinius sobolevi (Mukhina, 1970), and Merlinius tatrensis (Sabova, 1967). Tylenchorhynchus galeatus Litvinova, 1946 is with- drawn from the genus Merlinius. The following synonymies are made: Merlinius berberidis (Sethi and Swarup, 1968) is synonymized to M. hexagrammus (Sturhan, 1966); Ttjlenchorhynchus chonai Sethi and Swarup, 1968 is synonymized to T. triglyphus Seinhorst, 1963; Quinisulcius nilgiriensis (Seshadri et al., 1967) is synonymized to Q. acti (Hopper, 1959); and Tylenchorhynchus tener Erzhanova, 1964 is regarded a synonym of T. clarus Allen, 1955. The following are regarded in species inquirendae: TetylencJius dimidius Kirjanova, 1951; Aphelenchus dubius Steiner, 1914; and Fratylenchoides gadeai Arias-D, Jimenez-M. and Lopez-P., 1965.

Since my first compendium on Ttjlencho- proposed the new subfamily Merliniinae to ac- rhynchus (Tarjan, 1964), there have been a commodate Merlinius and formed two new number of new species proposed, two addi- genera within the Tylenchorhynchinae made tional keys prepared (Baqri and Jairajpuri, up of certain species formerly in Tylenchorhyn- 1970 and de Guiran, 1967), and new genera chus. The first of these is Uliginotylenchus proposed which are closely related to Tylen- with three incisures comprising an areolated chorhynchus (Siddiqi, 1970, 1971; Thorne and lateral field, female tail clavate to cylindroid Malek, 1968). In his first paper Siddiqi and with over 25 annules, and proximal end (1970) erected the new subfamily Tetylen- of gubernaculum bent dorsally. The second chinae to include the genus Tetylenchus and genus, Quinisulcius, has a nonareolated lateral proposed a new genus Merlinius to accommo- field with five incisures, distal flanges of date species of Tylenchorhynchus having six spicula are small-sized, and the proximal end incisures in the lateral field, a small trough- shaped nonprotrusible gubernaculum, and stout of gubernaculum directed dorsally. spicules with distal ends notched and without It is the object of the present study to large ventral flanges. Siddiqi's second paper evaluate the position of Tijlenchorhynchus and closely related genera, to determine member- 1 Florida Agricultural Experiment Stations Journal Series ship in such genera, and to present keys and a No. 4450. -Professor (Nematologist), University of Florida, IF AS, table of diagnostic data on species. The work Agricultural Research and Education Center, Lake Alfred, Florida 33850. presented was compiled only from publica-

Copyright © 2011, The Helminthological Society of Washington 124 PROCEEDINGS OF THE HELMINTHOLOGICAL SOCIETY tions and not from actual examination of present status of Trichotylenchus Whitehead, specimens. 1959 and Telotylenchus Siddiqi, 1960. Jairajpuri (1969) stated that Trichotylenchus differed Subfamily considerations from Telotylenchus only by a slightly narrower The Tylenchorhynchinae are, and have been, head, delicate spear, and three-incisured lateral a relatively homogenous group readily recog- field. In a later paper (1971), he presented nized by their cuticular markings, labial struc- a more detailed account corroborating and ture, stylet characteristics, esophagus structure, illustrating these differences. Jairajpuri re- sexual system arrangement, and tail shape. I garded Telotylenchus a junior synonym of see no practical value in assigning such closely Trichotylenchus. Seinhorst (1971) did not related genera as Tetylenchus, Tylenchorhyn- refer to the Jairajpuri references and regarded chus, and Merlinius to three different sub- the two genera as distinct, separated by stylet families, e.g., Tetylenchinae, Tylenchorhyn- appearance, shape of female tail, lateral field, chinae, and Merliniinae, respectively (Siddiqi, and shape of gubernaculum. He included 1971). Accordingly, I prefer to follow the ar- Trichotylenchus falciformis Whitehead, 1959; rangement suggested by Golden (1971) in T. rectangularis Netscher and Germani, 1969; recognizing one subfamily, the Tylenchorhyn- T. rhopalocercus (Seinhorst, 1963); T. palustris chinae, in which the new genus Merlinius (Merny and Germani, 1968); T. uliginosus (Siddiqi, 1970) can also be conveniently (Siddiqi, 1970); and T. papyrus (Siddiqi, accommodated. 1970) in the genus. He justified inclusion of some of these species, which in their original Previous departures descriptions do not all have the same type of basal portion of the esophagus, by suggesting In addition to the accurate list given by that the original authors misinterpreted the Baker (1962), Table 2 cites departures from true structure of the esophagus. Seinhorst's Tylenchorhynchus as given by various authors placement of these species thus results in a during the past two decades. genus in which: (a) two species (falciformis and rectangularis) have pronounced esopha- The genus Uliginotylenchus Siddiqi, 1971 geal overlaps, while four species do not; (b) This genus was characterized essentially as five species have delicate or slender stylets having an areolated lateral field with three in- while one (palustris) does not; (c) five species cisures, moderately sclerotized cephalic frame- have three-incisured lateral fields while one work, labial region not offset, spermatheca (rectangularis) has four incisures; and (d) offset, spicules with large-sized distal flanges, four species have somewhat clavate tails while gubernaculum with proximal portion directed two species (rectangularis and palustris) have dorsally, female tail with 25 or more annules cylindrical tails. Then, too, no mention is and broadly rounded, and occurring in wet made of Whitehead's (1959) original char- soil. The characters above appear to be valid acterization of Trichotylenchus as having a except for the cephalic framework which is head divided into four lobes and a distinctly invariably lightly, not moderately, sclerotized forked stylet. Because of these discrepancies for the five species included and for the offset and because I feel that the distinct and pro- nature of the spermatheca which is unknown nounced overlap of the esophagus on the in- for four of the five species. Of the species testine is of cardinal importance in identifying included, U. bifasciatus (Andrassy, 1961) ap- species of Trichotylenchus, I regard Siddiqi's pears to deviate most from the generic criteria (1971) assignment of species as the most in that: (a) its lip region is weakly offset, not logical. continuous; (b) its males do not show spicules It would be convenient to include the four with large-sized distal flanges; and (c) its remaining three-incisured "Tylenchorhynchus" type locality was in garden soil, not marshy or species in Uliginotylenchus, i.e., T. chonai, T. wet soil. divittatus, T. sculptus, and T. triglyphus. This Uliginotylenchus cannot be evaluated with- is not feasible principally because: (a) none out referring to the current controversy on the of the species have areolated lateral fields,

broadly rounded female tails with more than The generic diagnosis given by Golden (1971), 25 annules, or occur in wet soils, and (b) most patterned after previous diagnoses, is more of these species do not have gubernacula with inclusive. However, one of the principal char- proximal parts directed dorsally, spicules with acters of the genus, "cephalic framework large-sized distal flange, or lightly sclerotized present," becomes suspect when diagnostic cephalic frameworks. data on Tylenchorhynchus species is inspected (Table 1). Under the heading of "framework Key to Females of Uliginotylenchus* sclerotization" it can be seen that among 45 1. Tail with 24-35 annules, lip region with species, 13 have inconspicuous cephalic frame- 4-5 annules, tail terminus smooth ____ works, 6 have frameworks which were not ____ palustris (Merny and Germani, 1968) mentioned or depicted, while 15 have lightly Siddiqi, 1971 sclerotized frameworks. Tail with 42-56 annules, lip region with Siddiqi (1970) transferred Tylenchorynchus 5—8 annules, tail terminus annulated 2 galeatus Litvinova, 1946 to Merlinius. He later 2. T/ABW = 2.6-2.7, lip region 5-6 an- agreed (in litt.) that the species best be taken nules, lip region weakly offset out of Merlinius because it is reported to bifasciatus (Andrassy, 1961) have only four incisures in the lateral field. Siddiqi, 1971 Accordingly, it is returned to the genus T/ABW = 3.7-5.6, lip region with 6-8 Tylenchorhynchus. annules, lip region continuous 3 The only difference existing between the 3. Tail shape subcylindrical to slightly cla- description and figures for T. clarus Allen, vate, T/ABW = 5.6, tail with 42 an- 1955 and T. tener Erzhanova, 1964 is the b nules __ rhopalocercus (Seinhorst, 1963) ratio which is 4.0—5.0 and 5.3-9.8, respectively. Siddiqi, 1971 In almost all other important respects, e.g., Tail shape clavate, T/ABW < 5.6, tail stylet length, number of tail annules, tail—anal with 52-56 annules 4 body width ratio, head and tail shape, etc., 4. Stylet 14-16 ^ long, T/ABW = 4.6- differences are not evident. Therefore, T. 5.0, body 0.40-0.64 mm long tener Erzhanova, 1964 is regarded a junior uliginosus (Siddiqi, 1970) synonym of T. clarus Allen, 1955. Siddiqi, 1971 Tylenchorhynchus gadeai (Arias-D., Jimenez- Stylet 23-24 ^ long, T/ABW = 3.7- M., and L6pez-P., 1965) was originally de- 4.2, body 0.80-0.94 mm long scribed as a Pratylenchoides. Braun and Loof papyrus (Siddiqi, 1970) (1966) examined two type specimens and Siddiqi, 1971 concluded that the species belonged to Tylen- chorhynchus but stated that the original de- The genus Tylenchorhynchus Cobb, 1913 scription may have involved more than one species. Neither one of the accounts give the Because of the appearance of new closely number of incisures in the lateral field. Since related and more exactly defined genera, the description is lacking essential details and Tylenchorhynchus becomes the depository for since reasonable doubt exists as to its true species with four incisures in the lateral field, identity, the species is regarded as species as well as the four species with three incisures inquirenda. without areolation. The generic diagnosis Pertinent data and figures for T. triglyphus given by Siddiqi (1970) is inadequate because Seinhorst, 1963 and T. chonai Sethi and the generic criteria listed apply only to some Swarup, 1968 are almost identical. The only species, not all. For example, males are un- differences are claimed to be in head shape known for nine of the 46 species included; and stylet length (25-28 //, for chonai: 20- accordingly it is impractical to rely on precise 23 /j, for triglyphus). Inspection of the figures male characters as characterizing the genus. for each species fails to show any essential differences in head shape and a 2-ju, difference * The keys presented are based primarily on female characteristics except that male characters are also used in in stylet length ranges is not considered suf- some cases. Any abbreviations or symbols used are ex- plained at the end of Table 1. ficient difference for recognizing separate spe-

Copyright © 2011, The Helminthological Society of Washington 126 PROCEEDINGS OF THE HELMINTHOLOGICAL SOCIETY cies. Accordingly, T. chonai is regarded a 7. Tail terminus annulated 8 junior synonym of T. triglyphus Seinhorst, Tail terminus smooth 9 1963. 8. Stylet 24-28 p long, T/ABW = 1.9, Golden (1971) recently proposed the bino- V% = 46-53 men Tylenchorhynchus annulatus (Cassidy, lamelliferus (de Man, 1880) 1930) Golden, 1971 for Chitinotijlenchus Filipjev, 1936 annulatus after studying Cassidy's original Stylet 21 p long, T/ABW = 2.2-2.6, specimens. He stated, "This nematode is V% = 54-56 .. judithae Andrassy, 1962 clearly a Tylenchorhijnchus species closely re- 9. Stylet 16 ^ long, tail shape subcylin- lated to T. martini Fielding, 1956." Since drical, tail terminus hemispherical Golden did not further describe the species brevttineatus Williams, 1960 nor add it to its original description and since, Stylet 19—27 //, long, tail shape co- under Tylenchorhynchinae standards, the de- noid, tail terminus bluntly pointed 10 scription of the species is questionable, it is 10. Body with 16-20 longitudinal striae, best to regard it as species inquirenda as Slier tail with 53-55 annules, body (1970) suggested. 0.77-0.94 mm long microphasmis Loof, 1960 Key to Females of Tylenchorhynchus Body with 12 longitudinal striae, tail 1. Lateral field with 3 incisures 2 with 22-50 annules, body 0.54- Lateral field with 4 incisures 4 0.77 mm long 11 2. Cuticle of lateral fields extended on 11. Bursa distally recurved, female tail with 22 annules, spicules 24 ^ long female tail, stylet 16-17 ^ long, lip region offset „___ divittatus Siddiqi, 1961 phaseoli Sethi and Swarup, 1968 Cuticle of lateral fields normal on fe- Bursa continuing to end of tail, female tail, stylet 20-28 //, long, lip male tail with 30—50 annules, spic- region continuous 3 ules 27 ^ long sulcatus de Guiran, 1967 3. Tail with 9 annules, cephalic frame- 12. Tail terminus annulated 13 work heavily sclerotized, anterior Tail terminus smooth 23 surface of stylet knobs inclined 13. Stylet 27-38 /* long 14 anteriorly ____ sculptus Seinhorst, 1963 Stylet 17-24 p long 15 Tail with 13-15 annules, cephalic 14. Tail with 13-14 annules, T/ABW = framework lightly sclerotized, ante- 1.1, stylet 37-38 p long rior surface of stylet knobs inclined brevicaudatus Hopper, 1959 laterally _^__ triglyphus Seinhorst, 1963 Tail with 31-33 annules, T/ABW = (syn. T. chonai Sethi 2.5, stylet 27-30 ^ long and Swarup, 1968) magnicauda (Thorne, 1935) 4. Body with longitudinal striae 5 Filipjev, 1936 Body without longitudinal striae 12 15. Tail with 20-29 annules, lip region 5. Body with 29 longitudinal striae at with 4 annules 16 midbody, tail with 10 annules Tail with 31-66 annules, lip region claytoni Steiner, 1937 with 3 or 5—7 annules 17 Body with 20 or less longitudinal 16. Cuticular annulation irregular, lip striae, tail with 13 or more annules 6 region offset, anterior surface of 6. Lip region with 1-2 annules, tail with stylet knobs inclined posteriorly __ 13 annules, anterior surface of irregularis Wu, 1969 stylet knobs inclined anteriorly ____ Cuticular armulatiori coarse, lip region continuous, anterior surface of pachys Thorne and Malek, 1968 stylet knobs inclined anteriorly .... Lip region with 5—7 annules, tail with eremicolus Allen, 1955 22 or more annules, anterior sur- 17. Tail with 66 annules, anterior surface face of stylet knobs inclined poste- of stylet knobs inclined anteriorly riorly or laterally 7 canalis Thorne and Malek, 1968

Tail with 31-48 annules, anterior sur- offset, tail subcylindrical to conoid, face of stylet knobs inclined poste- tail terminus subhemispherical to riorly or laterally 18 bluntly pointed 28 18. Stylet 21-24 ^ long 19 28. Tail conoid, tail terminus bluntly Stylet 17-19 ^ long 20 pointed, body 0.65—0.99 mm long, 19. Tail subcylindrical, tail terminus sub- V% = 54-65 __ cylindricus Cobb, 1913 hemispherical, body 0.76-0.86 mm Tail subcylindrical, tail terminus sub- long bryobius Sturhan, 1966 hemispherical, body 0.50-0.62 mm Tail cylindrical, tail terminus hemi- long, V% = 52-54 spherical, body 0.98-1.40 mm long tarjani Andrassy 1969 maximus Allen, 1955 29. Lip region with only 2 annules 30 20. Tail shape clavate, lip region with 3 Lip region with 3 or more annules _„_ 31 annules, T/ABW = 3.8-4.0 30. Tail conoid, tail terminus hemispheri- davicauda Seinhorst, 1968 cal, T/ABW = 2.8, tail with 18- - (syn. clavicaudatus Seinhorst, 1963) 20 annules nudus Allen, 1955 Tail shape cylindrical to subcylindri- Tail subcylindrical, tail terminus sub- cal, lip region with 5-7 annules, hemispherical, T/ABW = 4.0, tail T/ABW = 2.0-3.2 21 with 29 annules 21. T/ABW = 2.0, spicule 30 p. long, lip delhiensis Chawla et al., 1968 region with 5 annules 31. Tail with 8-15 annules 32 huesingi Paetzold, 1958 Tail with 15-40 annules 34 T/ABW = 3.0-3.2, spicule 12-24 p 32. Tail cylindrical, tail terminus hemi- long, lip region with 7 annules .... 22 spherical, tail with 8—10 annules 22. Tail subcylindrical, tail terminus sub- georgiensis Eliashvili, 1971 hemispherical, tail with 46-48 an- Tail conical, tail terminus bluntly nules dubius (Biitschli, 1873) pointed, tail with 10-15 annules „ 33 Filipjev, 1936 33. Lip region offset, T/ABW = 2.2 .... Tail cylindrical, tail terminus hemi- latus Allen, 1955 spherical, tail with 35^43 annules Lip region continuous, T/ABW = 2.6 pawns Allen, 1955 clarus Allen, 1955 23. Tail with 40-58 annules 24 (syn. T. tener Tail with 8-40 annules 26 Erzhanova, 1964) 24. Tail with 40-45 annules, stylet 23 p. 34. Lip region with 3-4 annules 35 long, tail cylindrical, tail terminus Lip region with 5-7 annules 40 hemispherical 35. Stylet 11-14 p, long, tail with 28-32 robustus Thorne and Malek, 1968 annules .. ventrosignatus Tobar-J., 1969 Tail with 45-58 annules, stylet 28- Stylet 16-23 p long, tail with 15-33 39 p. long, tail subcylindrical or annules 36 conoid, tail terminus subhemi- 36. Lip region well set off, tail conoid, spherical 25 tail terminus bluntly pointed 25. Tail with 45-47 annules, stylet 31- brassicae Siddiqi, 1961 39 p long, T/ABW = 2.2 Lip region continuous or only weakly galeatus Litvinova, 1946 set off, tail cylindrical to subcylin- Tail with 51-58 annules, stylet 28-31 drical, tail terminus hemispherical p. long, T/ABW = 3.3-4.8 or subhemispherical 37 kegenicus Litvinova, 1946 37. Spermatheca and males not observed 26. Stylet 23-27 p. long 27 martini Fielding, 1956 Stylet 11-23 p. long 29 Spermathecae and males present .... 38 27. Tail with 17-23 annules, lip region 38. Stylet 20-23 p, long, spicule 22-25 p. continuous, tail cylindrical, tail ter- long, anterior surface of stylet minus hemispherical knobs inclined laterally silvaticus Ferris, 1963 agri Ferris, 1963 Tail with 14—16 annules, lip region Stylet 16-20 p, long, spicule 18-22 p.

long, anterior surface of stylet veloped bursa presumably is a sound character knobs inclined posteriorly 39 since it so appears in three species. Males for 39. Stylet 16-19 p long, tail terminus the remaining four species in the genus have subliemispherical to hemispherical not yet been discovered. The cephalic frame- mashhoodi Siddiqi and Basir, 1959 work is only moderately sclerotized in one Stylet 18—20 ^ long, tail terminus species, lightly sclerotized in three, incon- subliemispherical spicuous in two, and not referred to in one ewingi Hopper, 1959 species. I regard spicules in only one species 40. Lip region well offset, stylet 13-15 as bearing small-sized distal flanges; the illus- p, long goffarti Sturhan, 1966 trations of two species do not show flanged Lip region continuous or weakly set spicules while males have not been found in off, stylet 15-22 ^ long 41 four species. 41. Tail with 35-38 annules, T/ABW = The diagnostic data for Quinisulcius nilgi- 3.3 manubriatus Litvinova, 1946 riensis (Seshadri et al., 1967) and Q. acti Tail with 16-27 annules, T/ABW = (Hopper, 1959) closely parallel each other. 2.0-2.8 42 There are no apparent major differences. The 42. Stylet 21-22 p, long, cephalic frame- former species was said to differ from the work heavily sclerotized latter by the slope of the stylet, a slight esopha- ebriensis Seinhorst, 1963 geal overlap, and shape of the tail terminus. None of these differences are deemed of Stylet 15-18 p. long, cephalic frame- sufficient weight to justify acceptance of a work lightly to moderately sclero- separate species. Accordingly, Q. nilgiriensis tized 43 (Seshadri et al., 1967) is considered a syn- 43. Lateral field areolated, lip region set onym of Q. acti (Hopper, 1959). off, cephalic framework moderately The description for Tylenchorhynchus sclerotized .... aerolatus Tobar-J., 1970 acutoides Thorne and Malek, 1968 was ac- Lateral field not areolated, lip re- companied by illustrations incorrectly labeled. gion continuous, cephalic frame- Figure 16, C—F on page 43 of Thorne and work lightly sclerotized 44 Malek (1968) was identified as "T. paracutus." 44. Body 0.58-0.72 mm long, proximal This obviously was a lapsus calami; the nomen end of gubernaculum curved an- should be "T. acutoides." teriad almost 90° ._ striatus Allen, 1955 Body 0.42-0.63 mm long, proximal Key to Females of Ouinisulcius end of gubernaculum only slightly 1. Stylet 20-24 ^ long, lip region con- curved anteriad ._ contractus Loof, 1964 tinuous, tail with 49-56 annules goodeyi (Marinari, 1962) The genus Quinisulcius Siddiqi, 1971 Siddiqi, 1971 The salient characteristic of this genus was Stylet 15-19 /JL long, lip region offset, given as five incisures in a nonareolated lateral tail with 15-42 annules 2 field. An offset lip region, moderately sclero- 2. Lip region with 8 annules, tail with tized framework, ventrally arcuate female tail, 26-42 annules 3 well-developed bursa, spicules with small distal Lip region with 4-6 annules, tail with flanges, and protrusible gubernaculum with 15-23 annules 4 proximal end directed dorsally were given as 3. Tail with 42 annules, tail terminus additional characters. Of these characters, hemispherical, anterior surface of only a lateral field with five incisures is distinc- stylet knobs inclined laterally tive to all included species. The lateral field acti (Hopper, 1959) generally is nonareolated although Quinisulcius Siddiqi, 1971 cacti has an areolated lateral field anterior to (syn. nilgiriensis the metacorpus, but not posterior to it. The Seshadri et al, 1967) lip region is generally offset (exception: Q. Tail with 26 annules, tail terminus goodeyi Marinari, 1962) and the well-de- bluntly pointed, anterior surface of

stylet knobs inclined posteriorly cumbersome, erection of the genus Merlinius capitatus (Allen, 1955) is justifiable. Siddiqi, 1971 The following species are transferred into 4. Spermatheca and males present, body Merlinius: 0.80 mm long M. alboranensis (Tobar-Jimenez, 1970) _ acutoides (Thorne and Malek, 1968) comb. n. Siddiqi, 1971 M. brachycephalus (Litvinova, 1946) Spermatheca and males absent, body comb. n. 0.49-0.70 mm long 5 M. gaudialis (Izatullaeva, 1967) comb. n. 5. T/ABW = 2.1, anterior surface of M. parobscurus (Mulvey, 1969) comb n. stylet knobs inclined anteriorly „„ M. polonicus (Szczygiel, 1970) comb. n. acutus (Allen, 1955) M. sobolevi (Mukhina, 1970) comb. n. Siddiqi, 1971 M. tatrensis (Sabova, 1967) comb. n. T/ABW = 2.7-3.0, anterior surface Siddiqi (1970) questioned the transfer of of stylet knobs inclined laterally or brachycephalus to Merlinius. Litvinova (1946) posteriorly 6 depicted two tails for this species, one of which 6. Tail with 23 annules, lip region with certainly is not at variance with tails of some 6 annules, body 0.60-0.70 mm other species in the genus. Also, the head of long cacti (Chawla et al., 1968) the species, although unusually flat, bears af- Siddiqi, 1971 finities to other species in the genus. Transfer Tail with 15-22 annules, lip region of the species to Merlinius is logical. with 4-5 annules, body 0.49-0.63 The nomen Merlinius nanus (Allen, 1955) mm long curvus (Williams, 1960) Siddiqi, 1970 is not included in the list of Siddiqi, 1971 species given by Siddiqi (1970). However, he makes this combination in his illustrations and in a discussion of deirids within the same The genus Merlinius Siddiqi, 1970 paper. Perhaps the most important alteration oc- Siddiqi (1970) listed "M. dubius (Steiner, curring within the Tylenchorhynchinae has 1914) comb, n." The history of this species been the establishment of the genus Merlinius begins with Steiner (1914) describing a to accommodate those forms, previously in population of females from Switzerland as Tylenchorhynchus, that have six incisures in Aphelenchus dubius, having a wide lateral the lateral field. In the generic diagnosis field with longitudinal striations, a "Tylench"- presented by Siddiqi (1970), two additional type head, and a typical Aphelenchus-type characters in males are claimed to be definitive. tail. Goodey (1932) arbitrarily decided that These are: (a) distal end of spicules broadly a bisexual population which he found was con- rounded and notched, and (b) a nonprotrusi- specific with Steiner's species. Since Goodey ble gubernaculum. In personal correspondence regarded these as species of Anguillulina, and with Siddiqi, I pointed out that there are a Anguillulina dubius (Biitschli, 1873) pre- number of species reported without males empted use of Steiner's specific epithet, Goodey which were included in the genus and that gave the new name of Anguillulina macrura males were illustrated for some species with to the species. Allen (1955) examined topo- pointed spicules and/or protruding guber- type specimens and found that the species nacula. His response was logical in pointing was actually composed of a large form and a out that the salient and consistent generic small form. Wallace and Greet (1964) con- character is the six-incisured lateral field and curred and designated the small form as that some illustrations by authors may not be Tylenchorhynchus macrurus (Goodey, 1932) wholly accurate. Although there can be reluc- tance to accept a new genus based on only while the large form became T. icarus Wallace one character, viz. six incisures, the character and Greet, 1964. Finally, Siddiqi (1970) is consistent and easily recognizable. Since transferred both species to Merlinius. Since, the objective of such action is to make however, Tylenchorhynchus dubius (Biitschli, the unwieldly genus Tylenchorhynchus less 1873) remained in another genus, Steiner's

Copyright © 2011, The Helminthological Society of Washington 130 PROCEEDINGS OF THE HELMINTHOLOGICAL SOCIETY original specific name once again became of concrete criteria separating the species, M. valid and the species became Merlinius dubius berberidis (Sethi and Swamp, 1968) is re- (Steiner, 1914) Siddiqi, 1970. garded as a junior synonym of M. hexagrammus In thus reviewing the history of this species, (Sturhan, 1966). I fail to find any justification for Goodey (1932) to assume that his species from En- Key to Females of Merlinius gland was the same as Steiner's Swiss popula- 1. Body with longitudinal striae 2 tion. Even Allen (1955, p. 130) pointedly Body without longitudinal striae 11 assumed that Goodey's identification was cor- rect. Steiner's account lacks much essential 2. Tail with 43-60 annules 3 information and it is questionable whether his Tail with 21-40 annules 4 species again could be recognized. It is inter- 3. Tail with 58-60 annules, stylet 17-19 esting that Steiner (1920), Micoletzky (1922), /x, long, tail subcylindrical, tail ter- and Goffart (1930) transferred this species minus subhemispherical into genera that now are Helicotylenchus, hexincisus (Jairajpuri and Baqri, 1968) Tylenchorhijnchus, and Rotylenchus, respec- Siddiqi, 1970 tively. Accordingly, Aphelenchus dubius Tail with 43-46 annules, stylet 23- Steiner, 1914 [= Merlinius dubius (Steiner, 25 /x long, tail conical, tail termi- 1914) Siddiqi, 1970] is regarded as species nus bluntly pointed inquirenda, while Anguillulina macrura Goodey, koreanus Choi and Geraert, 1971 1932, which is adequately described and illus- 4. Body with 30-48 longitudinal striae trated, becomes Merlinius macrurus (Goodey, at midbody 5 1932) Siddiqi, 1970 because of its citation Body with 20-28 longitudinal striae in synonymy with M. dubius. at midbody 8 Merlinius sobolevi (Mukhina, 1970) comb, 5. Body with 48 longitudinal striae, tail n. was originally described as having 10-12 terminus annulated, stylet 18—20 ^ longitudinal striations in the lateral field and long tesselatus (Goodey, 1952) illustrated with 12 striations. It is sometimes Siddiqi, 1970 possible to detect faint lines between the Body with 30-36 longitudinal striae, striations in a lateral field. Assuming that this tail terminus smooth, stylet 20—24 was the case, M. sobolevi would then have the fji long 6 usual six longitudinal striations. This thesis 6. Lip region continuous, tail with 26- is supported by the observation of Geraert 27 annules, T/ABW = 3.7 (1966) concerning M. microdorus (Geraert, cylindricaudatus (Ivanova, 1968) 1966). He wrote, "The lateral field bears six Siddiqi, 1970 incisures but in quite a number of animals Lip region offset, tail with 22—24 an- I had the impression that additional incisures nules, T/ABW = 2.7-3.0 7 were present between the original six. Closer examination revealed rows of punctations be- 7. Tail terminus hemispherical, lip re- tween the incisures. . . ." Accordingly, M. sobo- gion with 5 annules, body 0.64- levi is judged to have six longitudinal lines in 0.73 mm long quadrifer (Andrassy, 1954) the lateral field. A comparison of the critical diagnostic data Siddiqi, 1970 between Merlinius hexagrammus (Sturban, Tail terminus bluntly pointed, lip re- 1966) and M. berberidis (Sethi and Swarup, gion with 6-7 annules, body 0.80- 1968) shows no differences, with the sole 0.90 mm long „ rugosus (Siddiqi, 1963) exception of body length (1.04—1.28 mm for Siddiqi, 1970 hexagrammus; 0.72-0.95 for berberidis). Sethi 8. Body with 20 longitudinal striae, and Swarup apparently were unaware of the body 0.55-0.58 mm long existence of Sturhan's species at the time they sobolevi (Mukhina, 1970) published. It has been amply shown in the comb. n. literature that temperature and host can affect Body with 24—28 longitudinal striae, body length of populations. With the absence body 0.63-1.04 mm long 9

9. T/ABW = 3.7, stylet 18-20 p. long 19. Stylet 12-15 p, long, lip region with lenorus (Brown, 1956) 7 annules, T/ABW = 3.8 Siddiqi, 1970 nanus (Allen, 1955) T/ABW = 1.9-2.3, stylet 20-24 p. Siddiqi, 1970 long 10 Stylet 16-18 p. long, lip region with 10. T/ABW = 1.9, stylet 24 p. long, tail 6 annules, T/ABW = 3.0 with 30-37 annules nothus (Allen, 1955) tartuensis (Krall, 1959) Siddiqi, 1970 Siddiqi, 1970 20. Tail with 65 annules, T/ABW = 4.5, T/ABW = 2.3, stylet 20-22 p. long, body 1.25-1.50 mm long tail with 21-22 annules ____ parobscurus (Mulvey, 1969) stegus (Thorne and Malek, 1968) comb. n. Siddiqi, 1970 Tail with 40-60 annules, T/ABW = 11. Tail terminus annulated 12 3.0-4.0, body 0.63-1.20 mm long .. 21 Tail terminus smooth 24 21. Lateral fields areolated in posterior 12. Tail terminus hemispherical, T/ABW part of body, lip region with 5 an- = 2.0-2.8 13 nules, spicules 19 p long Tail terminus bluntly pointed or sub- polonicus (Szczygiel, 1970) hemispherical, T/ABW = 3.0-4.5 17 comb. n. 13. Tail with 20-21 annules, lip region Lateral fields not areolated, lip region with 4-5 annules with 6-9 annules, spicules 27-30 ^ gatidialis (Izatullaeva, 1967) long 22 comb. n. 22. Lip region well offset, stylet 20—21 p Tail with 35-59 annules, lip region long laminatus (Wu, 1969) with 5-10 annules 14 Siddiqi, 1970 14. Lip region offset, stylet 21-23 p. Lip region slightly offset or continu- long, body 0.58-0.70 mm long .... ous, stylet 23-27 p. long 23 ^_ bogdanovikatjkovi (Kirjanova, 1941) 23. Spermatheca present, lip region with Siddiqi, 1970 7 annules, lip region continuous -___ Lip region continuous, stylet 24-42 obscums (Allen, 1955) /i long, body 0.83-1.96 mm long .... 15 Siddiqi, 1970 15. V% = 59-60, lip region with 5-6 Spermatheca not observed, lip re- annules, spicule 28-29 p. long gion with 8—9 annules, lip region socialis (Andrassy, 1962) slightly offset .... leptus (Allen, 1955) Siddiqi, 1970 Siddiqi, 1970 V% = 50-59, lip region with 8-10 24. Stylet 11-24 p, long 25 annules, spicule 39-40 //, long 16 Stylet 25-67 p. long 31 16. Stylet 25-34 p, long, tail with 39-47 25. Stylet 11-16 p. long, body 0.43-0.70 annules, body 0.83-1.19 mm long mm long 26 macrurus (Goodey, 1932) Stylet 20-24 p long, body 0.70-0.90 comb. n. mm long 28 Stylet 34-42 p, long, tail with 50-59 26. Stylet 11 /A long, tail with 26-33 an- annules, body 1.45—1.96 mm long nules, body 0.43-0.47 mm long .... icarus (Wallace and Greet, 1964) —.alboranensis (Tobar-Jimenez, 1970) Siddiqi, 1970 comb. n. 17. Stylet 12-18 p. long 18 Stylet 13-16 p. long, tail with 42-54 Stylet 20-27 p, long 20 annules, body 0.54-0.70 mm long 27 18. Lip region offset, cephalic framework 27. Lip region with 6 annules, spicule 22 heavily sclerotized p long, cephalic framework moder- undyferrus (Haque, 1967) ately sclerotized Siddiqi, 1970 brevidens (Allen, 1955) Lip region continuous, cephalic Siddiqi, 1970 framework lightly sclerotized 19 Lip region with 4 to 6 annules, spic-

ule 20-22 ju, long, cephalic frame- p. long macrodens (Allen, 1955) work lightly sclerotized Siddiqi, 1970 microdorus (Geraert, 1966) Stylet 25-38 p. long, male spicule 26- Siddiqi, 1970 34 p. long 36 28. Tail with 27-49 annules, T/ABW 36. Tail with 28-33 annules, lip region = 2.9—3.8, tail terminus bluntly with 8 annules .... affinis (Allen, 1955) pointed 29 Siddiqi, 1970 Tail with 20-23 annules, T/ABW = Tail with 36-52 annules, lip region 2.3-2.6, tail terminus hemispheri- with 6-7 annules 37 cal to subhemispherical 30 37. Lip region continuous 29. Lip region with 4 annules, lip region hexagrammus (Sturhan, 1966) continuous, T/ABW = 2.9 Siddiqi, 1970 bavaricus (Sturhan, 1966) (syn.: berberidis Sethi Siddiqi, 1970 and Swarup, 1968) Lip region with 6 annules, lip region Lip region well offset 38 offset, T/ABW = 3.8 38. Cephalic framework heavily sclero- obscurisulcatus (Andrassy, 1959) tized, tail with 36-42 annules Siddiqi, 1970 grandis (Allen, 1955) 30. V% = 58-59, stylet 23-24 p long, Siddiqi, 1970 anterior surface of stylet knobs in- Cephalic framework inconspicuous, clined posteriorly tail with 42-44 annules tatrensis (Sabova, 1967) lineatus (Allen, 1955) comb. n. Siddiqi, 1970 V% = 55, stylet 20-22 ^ long, anterior surface of stylet knobs inclined laterally varians (Thorne and Malek, 1968) Siddiqi, 1970 Thorne and Malek (1968) referred to the perioral disc, from which a slender stylet guide 31. Stylet 67 ^ long „ superbus (Allen, .1.955) extends back almost % the length of a very Siddiqi, 1970 Stylet 25-48 ^ long 32 slender stylet. They referred to an offset lip region and weakly developed cephalic frame- 32. Tail with 10-12 annules, T/ABW = work as distinguishing the genus. One species 1.1-1.5 was included, G. tenuidens Thorne and Malek, brachycephalus (Litvinova, 1946) 1968, from which the generic characters were comb. n. obtained. Tail with 28-58 annules, T/ABW = Dr. M. R. Sauer kindly let me examine what 2.3-3.6 33 appears to be a new species of Geocenamus 33. Lip region with 9 annules, T/ABW from Australia and which clearly shows the = 3.4-3.6 34 generic characters listed above. He further Lip region with 6-8 annules, T/ABW influenced me to reexamine the characters of = 2.0-3.0 35 Tijlenchorhynchus arcticus Mulvey, 1969 which 34. Tail terminus hemispherical, tail with he and R. H. Mulvey conclude belongs to 54—58 annules, cephalic framework Geocenamus. I concur and regard the taxon heavily sclerotized as Geocenamus arcticus (Mulvey, 1969) alpinus (Allen, 1955) comb. n. Siddiqi, 1970 The original description and illustrations for Tail terminus bluntly pointed, tail Tylenchorhynchus longus Wu, 1969 fulfill all with 49—55 annules, cephalic of the generic criteria listed above, except for framework sclerotization inconspic- reference to a slender stylet guide. Knobloch uous conicus (Allen, 1955) (1971) emended the description of the species Siddiqi, 1970 principally to include longitudinal striations on 35. Stylet 43-48 p. long, male spicule 37 the labial region and body, and enlarged

Copyright © 2011, The Helminthological Society of Washington OF WASHINGTON, VOLUME 40, NUMBER 1, JANUARY 1973 133 phasmids. However, she pointed out simi- The genus Tetylenchus Filipjev, 1936 larities of the species to Tylenchorhynchus Filipjev (1936) characterized this genus as mamillatus Tobar-Jimenez, 1966, which has be- "Head without chitinization, cuticle finely come Scutylenchus mamillatus (Tobar-Jimenez, striated (with some exceptions), esophagus 1966) Jairajpuri, 1971. Scutylenchus Jairajpuri, tylenchoid, ovaries double, spear of moderate 1971 was based on characters of S. mamillatus size, males unknown." He made T. tennis which included: (a) a subdigitate tail shape, (Micoletzky, 1922) the type and included T. (b) enlarged scutella-like phasmids, (c) areo- clavicaudatus (Micoletzky, 1922) and T. lated lateral field, and (d) sloping stylet knobs. granulosus (Cobb, 1893). Tetylenchus clavi- Admittedly, similarities between S. mamillatus caudatus (Micoletzky, 192.1.) was transferred and T. longus are noteworthy, viz.: (a) dou- to Psilenchus in 1949 by Thorne (1949) and ble-layered cuticle, (b) enlarged phasmids, later regarded as species inquirenda by and (c) tesselated nature of cuticle due to Jairajpuri (1966). Tetylenchus granulosus longitudinal striae. However, the dissimilarities (Cobb, 1893) was either overlooked or ignored between the two species are even more im- by authors until Slier (1968) placed it in syn- portant, i.e., (a) occurrence of perioral disc in onymy with Radopholus similis (Cobb, 1893) T. longus, (b) stylet long and slender for T. Thorne, 1949. Thus, the only remaining rep- longus, short and stout for S. mamillatus, and resentative to the genus is the type T. tennis (c) cephalic sclerotization weak for T. longus (Micoletzky, 1922). Micoletzky described T. and moderately strong for S. mamillatus. The tennis from one female. He felt that the long presence of a prominent perioral disc and per- esophagus (b = 3.8), clear annulation of the haps also the slender stylet in T. longus relate cuticle, delicate weakly knobbed stylet, am- the species more cogently to Geocenamus. phidelphic sexual system, and conical tail with Siddiqi (in litt.) stated that T. longus could minutely rounded terminus characterized the not be accommodated in Merlinius because of species. He depicted a head with delicate the conspicuous labial disc which he used to stylet showing what one could interpret as a separate Geocenamus from Merlinius. Accord- moderately sclerotized framework. Hence, it ingly, the species is designated Geocenamus is seen that Filipjev's definition of Tetylenchus longus (Wu, 1969) comb. n. based on Micoletzky's one female specimen deviates in regard to head sclerotization, cutic- Key to Females of Geocenamus ular annulation, and stylet size. Thorne 1. Stylet 53-65 ^ long, T/ABW = 2.0- (1949) emended the generic diagnosis. He 2.2 longus (Wu, 1969) diagnosed the genus as without sclerotized comb. n. cephalic framework, stylet with or without Stylet 27-38 p. long, T/ABW = 2.9- basal knobs, tails tapering to acute or subacuate 3.0 2 terminus, deirids and phasmids present, bursa subcaudal extending almost to terminus, etc. 2. Stylet 27 ^ long, body 0.83 mm long He placed four species in the genus, T. tennis, .___ tenuidens Thorne and Malek, 1968 and three new species. Stylet 34-38 p long, body 1.05-1.27 In view of the foregoing historical analysis, mm long arcticus (Mulvey, 1969) it appears that the genus, as it is now recog- comb. n. nized, differs considerably from the characters Micoletzky (1922) ascribed to T. tennis. It is The genus Nagelus Thorne and Malek, unfortunate that the genus is based on a type 1968 species described from only one female. Yet The asymmetrical fine striation of the lip it also becomes clear that Tetylenchus is de- region, inconspicuous cephalic framework, an- fined by some characters which are not valid, gular stylet knobs, attachment of stylet pro- e.g., the absence of cephalic sclerotization. An tractor muscles and presence of epiptygma at examination of illustrations and specimens of the vulva characterizes this genus. It has only T. joctus reveals the presence of a cephalic one species, N. aberrans Thorne and Malek, framework, admittedly very lightly sclerotized 1968. in some cases, but nonetheless there. This is

Copyright © 2011, The Helminthological Society of Washington 134 PROCEEDINGS OF THE HELMINTHOLOGICAL SOCIETY similar to the rest of the Tylenchorhynchinae 6. Stylet 12 ^ long, tail with 67 annules, for which the intensity of cephalic sclerotiza- lip region with 4-5 annules tion will vary within genera. Hence, the most productus Thorne, 1949 important single characterization of Tetylen- Stylet 15 /jL long, tail with 42 annules, chus would be the presence of an acute or sub- lip region with 7 annules acute female tail with finely rounded terminus. joctus Thorne, 1949 This character is unique among the genera in the subfamily. In addition, the cephalic Based on the foregoing data, the Tylencho- framework can be regarded as inconspicuous rhynchinae can be regarded as made up of to lightly sclerotized and never heavily sclero- eight genera. tized. In all other respects, Tetylenchus appears to be close to Tylenchorhynchus and Key to Tylenchorhynchinae Eliava, 1964 Merlinius. 1. Female tail subdigitate, phasmids en- Tylenchorhynchus aduncus de Guiran, 1967, larged ____ Scutylenchus Jairajpuri, 1971 because of the tail shape of both sexes, tail Female tail not subdigitate, phasmids terminus shape, and light cephalic sclerotiza- normal 2 tion shown in the figures, fulfills the criteria 2. Female tail acute or subacute, termi- stated above and becomes Tetylenchus aduncus nus finely rounded (de Guiran) comb. n. Tetylenchus Filipjev, 1936 Tetylenchus dimidius Kirjanova, 1951 was Female tail bluntly conical to cylin- thought to be a Tylenchorhijnchus by Loof drical, terminus bluntly pointed to (1959). Merny (1964) also did not consider hemispherical 3 the species a true Tetylenchus. Both authors 3. Stylet knobs characteristically angu- felt that the cylindrical tail and subhemi- lar, protractor muscles of stylet at- spherical terminus precluded its placement in tached to inner wall of labial cavity, Tetylenchus; I concur. Accordingly Tetylen- epiptygma present at vulva chus dimidius Kirjanova, 1951 is regarded as Nagelus Thorne and Malek, 1968 species inquirenda. Stylet knobs rounded, protractor muscles of stylet attached to base of Key to Females of Tetylenchus cephalic framework, vulva seldom with epiptygma 4 1. Stylet without knobs abulbosus Thorne, 1949 4. Labial region with prominent peri- Stylet with knobs 2 oral disc, stylet characteristically slender 2. T/ABW = 5.7, V ratio 42 _ Geocenamus Thorne and Malek, 1968 tennis (Micoletzky, 1922) Labial region without perioral disc, T/ABW = 2.5-3.8, 'a' ratio 18-33.. 3 stylet moderately robust to heavy 5 3. Lateral field with 3-4 incisures 4 5. Lateral field with 3-4 incisures 6 Lateral field with 6 incisures 5 Lateral field with 5-6 incisures 7 4. Lip region offset, tail with 19-27 an- 6. Lateral field areolated, with 3 inci- nules, lateral field with 4 incisures sures ____ Uliginotylenchus Siddiqi, 1971 aduncus (de Guiran, 1967) Lateral field usually with 4 incisures, comb. n. never areolated when with 3 inci- Lip region continuous, tail with 17 sures ____ Tylenchorhynchus Cobb, 1913 annules, lateral field with 3 incisures annulatus Merny, 1964 7. Lateral field with 5 incisures Quinisulcius Siddiqi, 1971 5. Stylet 29 ^ long, tail with 35 annules, Lateral field with 6 incisures spicule 34 ^ long Merlinius Siddiqi, 1970 curiosus Wilski, 1964 Stylet 12-15 ^ long, tail with 42-67 Table 1 is a compendium of morphological annules, spicule 19-21 ^ long 6 measurements and other information of diag-

SMO SMO

TAIL SHAPES TAIL TIP SHAPES

CYL= CYLINDRICAL HEM= HEMISPHERICAL SCL=SUBCYLINDRICAL SHM = SUBHEMISPHERICAL CON--CONOID BLP--BLUNTLY POINTED ANN CLA= CLAVATE TAIL TIP ANNULATION ANN--ANNULATED SMO--SMOOTH Figure 1. Tylenchorhynchinae tail shapes and code designations used in Table 1. nostic value on the species within each genus, 1. Figure 1 is a diagrammatic explanation of as covered in the preceding keys. The table the tail codes used in Table 1. is based on data concerning females, except for the last two columns which refer to male Acknowledgments characters. When specific data were not presented in the text of the original publica- Miss Janell Kay Haglund, laboratory aide, tion, these were obtained from the accompany- contributed significantly to this work in pre- ing drawings, if possible, e.g., spicule length, paring tables of diagnostic data and by con- T/ABW (tail divided by body width at the scientiously reviewing the text for mistakes. anus), number of tail or head annules, etc. Dr. E. Krall amiably supplied photocopies of If certain information on a species was un- Russian references which were unavailable to known, a dash (—) was inserted in the table. me. Drs. M. R. Siddiqi, A. M. Golden, S. A. An explanation of the abbreviations and Slier, and R. P. Esser kindly reviewed the code symbols used appears at the end of Table manuscript.

Stylet Lateral No. No. Spicule Gubern. Uliginotylenchus Length Labial Labial Frm. length Sty. inci - long. tail Tail Tail length length Siddiqi, 197 1 in mm a b c V% region annules scl. in /t cli. sures striae annules shape terminus T/ABW in ft. in fi bifasciatus 0.65 - 26-3 1 5.6-5. 9 13-1 7 51-5 2 OF F 5- 6 LSC 19-2 0 PO S 3 42-4 6 SCL HEM-ANN 2.6-2. 7 25 12 ( Andrassy, 1961 ) 0.73 palustris 0.46 - 27-3 4 4.1-5. 8 12-1 6 52-5 9 CN T 4- 5 LSC 14-1 7 LA T 3 24-3 5 SCL HEM-SMO 2.7-3. 9 18-2 2 9-1 1 ( Merny and Ger- 0.62 mani, 1968 ) papyrus 0.80 - 37-4 3 5.5-6. 6 13-1 6 52-5 5 CN T 7- 8 LSC 23-2 4 PO S 3 56 CLA HEM-ANN 3.7-4. 2 26-2 9 13 (Siddiqi, Copyright © HelminthologicalThe 2011,of CopyrightSociety© Washingto 1970) 0.94 rhopalocercus 0.62 - 38-4 6 6. 5 11-1 3 49-5 5 CN T 7- 8 LSC 17-1 9 PO S 3 42 SCL HEM-ANN 5. 6 2 1 8 (Seinhorst, 1963 ) 0.81 uliginosus 0.40 - 30-3 7 4.4-5. 6 11-1 3 52-5 8 CN T 6- 7 LSC 14-1 6 LA T 3 52 CLA SHM-ANN 4.6-5. 0 19-2 1 10-1 1 (Siddiqi, 1970) 0.64

Tijlenchorhynchus Cobb, 191 3 acrolatus 0.54 - 26-3 1 4.7-6. 0 15-1 8 54-5 9 OF F 5- 6 MSC 15-1 7 PO S 4 16-2 2 SCL BLP-SMO 2.0-3. 0 19-2 2 10-1 2 Tobar- Jimenez, 0.70 1970 agri 0.66 - 28-3 3 4.7-5. 5 15-2 1 55-5 8 OF F 4 INC 20-2 3 LA T 4 18-2 6 SCL HEM-SMO 2.6 22-2 5 13-1 4 Ferris, 196 3 0.77 • brassicae 0.58 - 26-3 5 5.0-6. 0 14-1 7 52-5 8 OF F 4 LSC 16-1 7 LA T 4 18-3 3 CON BLP-SMO 2.5-3. 0 18-2 1 9-1 1 Siddiqi, 196 1 0.72 § brevicaudatus 1.28 - 34-3 6 6.7-6. 9 34-4 4 57-5 9 CN T 8 HSC 37-3 8 LA T 4 13-1 4 SCL HEM-ANN 1. 1 32 13 oO Hopper, 195 9 1.37 w brevilineatus 0.56 - 34 5.6 14 53-5 6 OF F 5- 6 LSC 16 PO S 4 8-12 ? 40-4 1 SCL HEM-SMO 2. 9 23 12 H Williams, 196 0 0.65 CE R O bnjobius 0.76 - 31-3 7 5.8-6. 6 14 52 CNT 5- 6 MSC 21-2 4 PO S 4 35-4 5 SCL SHM-ANN 2.5-3. 5 23-2 6 13 2, O Sturhan, 196 6 0.86 en canalis 1.00 39 7. 1 13 53 OF F 5- 6 MSC 20 AN T 4 66 SCL HEM-ANN 2. 8 25 13 Thome and Malek, O 1968 clarus 0.49 - 28-3 3 4.0-5. 0 16-2 0 57-6 1 CN T 5 LSC 15-1 7 AN T 4 10-1 5 CON BLP-SMO 2.6 p p ffit-H Allen, 195 5 0.61 W

clavicauda 0.54 - 31-4 3 4.5-5. 9 12-1 8 53-5 9 CN T 3 LSC 18-1 9 PO S 4 3 1 CLA HEM-ANN 3.8-4. 0 19 12 "T1 n Seinhorst, 196 8 0.72 w claijtoni 0.64 - 24-2 5 5,0-6. 2 18-1 9 55-5 7 OF F 3 ? 20 PO S 4 29 10 CON BLP-SMO 1. 7 25 15 t-1 Steiner, 193 7 0.73 MI D s contractus 0.42 - 22-3 0 3.8-5. 0 13-1 5 56-6 0 CNT 5- 6 LSC 16-1 8 LA T 4 20-2 3 CON SHM-SMO 2. 5 17-1 8 8- 9 2 Loof, 196 4 0.63 H ffi cylindriciis 0.65 - 28-3 5 4.2-6. 0 13-2 0 54-6 5 OF F 5 MSC 24-2 7 AN T 4 15-1 6 CON BLP-SMO 2. 5 25 16 O Cobb, 191 3 0.99 delhiensis 0.60 - 24-3 3 5.0-6. 0 13-1 8 47-4 8 CN T 2 INC 14-1 6 LA T 4 29 SCL SHM-SMO 4. 0 p p 8 Chawla et al., 196 8 0.70 o divittatus 0.55 - 32-3 8 5.4-6. 4 16-1 9 53-5 5 OF F 5 INC 16-1 7 LA T 3 21 SCL 'HEM'-SMO 2. 8 17 8 Siddiqi, 196 1 0.72 r dubius 0.62 - 30-3 5 5.0-6. 0 13-1 6 54-5 7 OF F 7 LSC 18-1 9 PO S 4 46-4 8 SCL SHM-ANN 3. 2 24 12 VI (Biitschli, 1873 ) 0.78 O Q ebriensis 0.52 - 23-2 9 4.9-5. 4 14-1 6 54-5 6 CN T 5 HSC 21-2 2 PO S 4 25 CON BLP-SMO 2. 6 21 10 Seinhorst, 196 3 0.59 H K< I Table 1 cont'd.

Stylet Lateral No. No. Spicule Gubern. Length Labial Labial Frm. length Sty. inci- long. tail Tail Tail length length in mm a b c V% region annules scl. in n cli. sures striae annules shape terminus T/ABW in /a in ILL eremicolus 0.70 - 28-3 0 4.5-5. 8 14-1 6 55 CN T 4 MSC 19-2 0 ANT 4 27-2 9 CON BLP-ANN 2.7 22 9 O Allen, 195 5 0.76 oH etcingi 0.55 - 30-3 5 4.3-5. 5 13-1 6 53-5 9 CN T 3 INC 18-2 0 POS 4 15-1 9 SCL SHM-SMO 2.5 22 12 Hopper, 195 9 0.75 galeatus 1.68 31 7. 3 19 54 CNT 10-1 3 ? 31-3 9 POS 4 45-4 7 CON SHM-SMO 2.2 51-53 ? 13-1 8 o Litvinova, 194 6 r georgiensis 0.57 - 29-3 5 4.3-5. 0 19-2 3 57-6 1 OF F 5- 6 ? 19-2 1 POS 4 8-1 0 CYL HEM-SMO 2.4 p ? d Eliashvili, 197 1 0.69 ft

Copyright © HelminthologicalThe 2011,of CopyrightSociety© Washingto goffarti 0.54 - 29-3 7 4.9-6. 0 13-2 0 52-5 7 OF F 6- 7 LSC 13-1 5 POS 4 23-4 0 SCL SHM-SMO 2.5-3. 5 21-2 3 11-1 3 Sturhan, 196 6 0.69 o tmesingi 0.78 - 26-31 5.0-5. 9 15-2 1 53-5 5 CN T 5 ? 18-1 9 POS 4 32-3 6 CYL HEM-ANN 2.0 30 14-1 5 Paetzold, 195 8 0.92 d irregularis 0.69 - 26-3 3 4.8-5. 8 15-1 9 55-5 9 OF F 4 LSC 19-2 1 POS 4 20-2 6 CON BLP-ANN 2.0-2. 5 26-2 9 16-1 8 Wu, 196 9 0.83 w judithae 0.88 - 33 6.7-7. 2 17-2 3 54-5 6 OF F 6- 7 LSC 21 LAT 4 14 36-4 0 CON SHM-ANN 2.2—2.6 p p H Andrassy, 196 2 0.97 kegenicus 0.79 - 25-3 5 4.3-6. 8 10-1 4 42-5 5 CNT 7-1 0 ? 28-3 1 POS 4 51-5 8 SCL SHM-SMO 3.3-4. 8 26-3 1 9 -1"1 Litvinova, 194 6 1.1 7 lamelifents 0.86 - 25-3 1 4.3-6. 8 16-2 4 46-5 3 CNT 6 INC 24-2 8 LAT 4 14CER 41-4 5 CON BLP-ANN 1.9 31 16 Z (deMan, 1880 ) 1.10 16 MID d latus 0.58 - 30-4 0 4.9-5. 4 18-2 2 56-6 1 OF F 6 LSC 16-1 7 ANT 4 14-1 5 CON BLP-SMO 2.2 p p fe Allen, 195 5 0.70 >< magnicauda 0.79 - 23-3 2 4.4-6. 0 13-1 9 56-6 2 CNT 6 HSC 27-3 0 ANT 4 31-3 3 CYL HEM-ANN 2.5 p ? ,_, cc (Thorne, 1935 ) 1.00 p manubriatus 0.75 29 5.4 15 55 CN T 6 ? 18 POS 4 35-3 8 SCL HEM-SMO 3. 3 p w Litvinova, 194 6 p martini 0.75 31 5. 0 14 54 OF F 3 ? 19 LAT 4 22-2 7 SCL HEM-SMO 3.7 p Fielding, 195 6 mashhoodi 0.49 - 26-3 7 4.4-5. 8 13-1 9 50-5 9 CN T 3- 4 LSC 16-1 9 POS 4 15-3 0 CYL SHM-SMO 2.5-4. 0 18-2 2 10-1 5 Siddiqi and Basir, 0.76 1959 maximus 0.98 - 37-4 7 5.4-8. 1 16-2 0 47-5 4 CNT 7 LSC 21-2 4 POS 4 38-4 1 CYL HEM-ANN 2.6 p p Allen, 195 5 1.40 microphasmis 0.77 - 31-3 9 5.2-6. 1 15-1 8 49-5 5 OF F 6- 7 INC 24-2 7 POS 4 16-2 0 53-5 5 CON BLP-SMO 3.0 29 14 Loot, 196 0 0.94

n nudus 0.71 30 4.1-4. 8 14-1 5 54-5 6 CNT 2 INC 19-2 3 ANT 18-2 0 CON HEM-SMO 2.8 23 13 Allen, 195 5 4 pachys 0.63 24 5.0 16 57 CNT 1- 2 LSC 13-1 5 ANT 16 1 3 CON BLP-SMO 2.4 23 15 Thorne and Malek, 4 MID 1968 parvus 0.65 - 25-3 0 4.8-5. 6 13-1 6 52-5 7 CNT 7 INC 17-1 8 LAT 4 35-4 3 CYL HEM-ANN 3.0 12 5 Allen, 195 5 0.72 phaseoli 0.61 - 28-3 5 4.0-5. 4 17-2 0 54-5 8 OF F 6 LSC 19-2 6 POS 4 12 22 CON BLP-SMO 2.3 24 12 Sethi and Swarup, 0.77 1968 robustus 1.00 31 7 15 52 CNT p MSC 23 ANT 4 40-4 5 CYL HEM-SMO 3.5 27 15 Thorne and Malek,

1968 h-*

F O > WASHI Tabl e 1 cont'd.

Stylet Latera l No. No. Spicule Gubern . Length Labia l Labia l Frm . lengt h Sty. inci - long. tai l Tai l Tail length lengt h in mm a b c V% region annules scl. in fi cli. sures striae annules shape terminus T/AB W in ft in IL sculptus 0.54- 23-2 5 4.3-4. 7 18-1 9 56-5 7 CN T 2- 3 HSC 22 ANT 3 9 SCL SHM-SMO 2.3 22 12 Seinhorst, 196 3 0.59 silvaticus 0.80 - 24-3 3 5.5-6. 3 18-2 3 53-5 8 CN T 4 INC 23-2 6 LAT 4 17-2 3 CYL HEM-SMO 2.5 29 18 Ferris, 196 3 1.00 striding 0.58 - 29-3 4 5.0-5. 3 13-1 6 55-5 7 CN T 5 LSC 16-1 7 ANT 4 20-2 7 SCL SHM-SMO 2.8 20 12 Allen, 195 5 0.72

Copyright © HelminthologicalThe 2011,of CopyrightSociety© Washingto sulcatus 0.54 - 27-3 7 5.1-5. 7 14-1 7 52-5 6 OF F 6- 7 INC 19-2 2 POS 4 12 30-5 0 CON BLP-SMO 2.5-3. 3 27 15 de Guiran, 196 7 0.71 tarjani 0.50 - 24-2 7 4.7-5. 5 16-1 8 52-5 4 OF F 4- 5 INC 24-2 5 POS 4 14-1 5 SCL SHM-SMO 2.7-2. 8 26 15 Andrassy, 196 9 0.62 triglyphu$ 0.58 - 24-3 0 4.6-5. 6 16-1 9 55-5 8 CN T 4 LSC 20-2 3 LAT 3 13-1 5 CON BLP-SMO 2. 5 22 10 Seinhorst , 196 3 0.68 ventrosignatus 0.45 - 28-3 6 3.9-6. 0 13-1 6 53-5 7 OF F 4 INC 11-1 4 POS 4 28-3 2 SCL BLP-SMO 2.6-3. 2 20-21 11 Tobar- Jimenez , 0.62 1969

Quinisulcius Siddiqi, 197 1 acti 0.65 - 30-3 6 4.6-4. 9 15-1 6 55-5 7 OF F 8 INC 17 LAT 5 — 24 CON HEM-SMO 3.0 p p . (Hopper, 1959 ) 0.71 acutoides 0.80 35 5. 4 20 57 OF F 4- 5 MSC 18 ANT 5 18 CON BLP-SMO 2.8 20 10 Thom e and Malek, a 1968) acutus 0.63 - 31-3 8 4.8-5. 3 17-2 2 56-5 7 OF F 6 LSC 16-1 7 ANT 5 17 CON BLP-SMO 2. 1 p p w (Allen, 0.70 1955 ) p a cacti 0.60 - 25-3 3 4.0-6. 0 14-1 9 54-5 9 OF F 6 ? 15-1 9 LAT 5 23 SCL SHM-SMO 3.0 p ( Chawla et al., 0.70 o 1968) C/3 capitatus 0.63 - 30-3 8 5.0-5. 8 12-1 7 51-5 8 OF F 8 LSC 16-1 8 POS 5 26 CON BLP-SMO 2.8 19 7 o (Allen, 1955 ) 0.85 ^ curvus 0.49 - 29-3 7 4.6-5. 5 15-1 8 52-5 7 OF F 4- 5 INC 17 POS 5 15-2 3 CON BLP-SMO 2.7 p p H (Williams, 1960 ) 0.63 B goodeyi 0.51 - 24-2 9 4.4-5. 4 14-1 6 52-5 6 CN T 6- 7 LSC 20-2 4 POS 5 49-5 6 CON HEM-ANN 2.8-3. 2 25-2 9 8-1 0 n (Marinari, 1962 ) 0.81 K Mr Merlinius § Siddiqi, 197 0 § H affinis 0.80 - 30-3 3 4.2-5. 0 14-1 5 55-5 8 CN T 8 HSC 26-2 7 POS 6 28-3 3 SCL SHM-SMO 2.3 27 9 K (Allen, 1955 ) 0.90 O alboranensis 0.43 - 25-3 0 4.5-5. 3 12-1 5 58-6 1 OF F 6 LSC 11 POS 6 26-3 3 SCL SHM-SMO 2.4-2. 8 17-20 5- 7 r ( Tobar - Jimenez, 0.47 1970 ) n.c. o ulpinus 0.83 - 26-2 7 5.0-5. 3 11-1 3 52-5 4 OF F 9 HSC 39-4 2 POS 6 54-5 8 SCL HEM-SMO 3.6 p p (Allen, 1955 ) 0.95 ^r bavaricus 0.75 28 5.2 13 57 CNT 4 LSC 21 POS 6 49 SCL BLP-SMO 2.9 p P Crt ( Sturhan, 196 6 ) O bogdanovikatjkovi 0.58 - 23-2 7 4.0-5. 0 13-1 4 54-5 8 OF F 7-9? MSC 21-2 3 POS 6 47? CON HEM-ANN 2.8 p p (Kirjanova, 1941 ) 0.70 H Table 1 cont'd.

Style t Latera l No . No . Spicul e Gubern . Lengt h Labia l Labia l Frm . lengt h Sty . inci - long . tai l Tai l Tai l lengt h lengt h in m m a b c V % regio n annule s sci . in /i cli. sure s stria e annule s shap e terminu s T/AB W in fa in fi ^: brachycephalus 0.66 - 26-2 8 4.4-5. 4 35-4 2 52-6 9 CN T 5- 7 HS C 28-3 0 PO S 6 10-1 2 CON BLP-SM O 1.1-1. 5 37 11 O (Litvinova , 1946 ) 0.7 7 g n.c. 2 brevidens 0.54 - 23-2 7 4.2-5. 2 11-1 3 52-5 8 CN T 6 MSC 14-1 6 PO S 6 42-4 9 SC L HEM-SM O 2. 8 15 10 (Allen , 1955 ) 0.6 9 conicus 0.78 - 31-3 9 4.4-5. 8 11-1 4 52-5 6 OF F 9 IN C 40-4 5 PO S 6 49-5 5 CO N BLP-SM O 3. 4 32 9 r (Allen , 1955 ) 1.10 d^ cylindricaiidatus 0.77 - 28-3 2 5.0-5. 8 14-1 6 57-6 0 CN T 7 LS C 22-2 4 PO S 6 30-3 4 26-2 7 SCL SHM-SM O 3. 7 p p 2 (Ivanova , 1968 ) 0.9 0 MID M Copyright © HelminthologicalThe 2011,of CopyrightSociety© Washingto gaudialis 0.66 - 23-2 6 4.7-6. 0 17-2 2 55-6 0 CN T 4- 5 MSC 22—25 POS 6 20-2 1 SC L HEM-AN N 2.0-2. 4 34-3 6 11-1 4 o ( Izatullaeva , 0.7 0 1957) n.c . 2 d grandis 0.96 - 34-4 1 5.0-6. 4 13-1 6 50-5 4 OF F 6 HS C 26-3 0 PO S 6 36-4 2 SC L SHM-SM O 2. 8 30 9 *-* (Allen , 1955 ) S 1.1 1 CO hexagrammus 1.04 - 27-3 4 5.2-6. 4 15-2 0 51-5 6 CN T 6- 7 HS C 33-3 6 PO S 6 39-5 2 CO N BLP-SM O 2.0-3. 0 32-3 4 11-1 2 H Sturhan , 196 6 1.28 tfl hexincisis 0.85 - 32-3 9 6.0-8. 2 18-2 3 52-5 5 OF F 7 IN C 17-1 9 POS 6 16 58-6 0 SCL SHM-AN N 3.0-4. 0 28-3 1 8- 9 ,M ( Jairajpur i and 1.08 MI D <-H Baqri , 1968 ) 2! teams 1.45 - 29-3 4 5.9-6. 9 19-2 5 50-5 7 CN T 8 HS C 34-4 2 PO S 6 50-5 9 CYL HEM-AN N 2. 2 40 12 d ( Wallac e and 1.96 >• Greet , 1964 ) 3 koreanus 0.75 - 30-4 4 5.1-6. 1 12-1 6 53-6 0 OF F 6 LS C 23-2 5 AN T 6 15-1 7 43-4 6 CO N BLP-AN N 3.4-3. 8 p p CO Cho i an d Geraert , 0.9 2 <1 197 1 CO laminatus 0.80 - 31-3 9 6.0-7. 3 13-1 6 52-5 5 OF F 6- 8 LS C 20-2 1 PO S 6 50 CO N BLP-AN N 3.0-3. 9 27-3 0 9-1 1 (Wu, 1969) 1.20 * lenorus 0.63 - 22-2 9 5.2-6. 7 12-1 5 52-5 8 OF F 6 LS C 18-2 0 PO S 24 28-3 0 CO N SHM-SM O 3. 7 27 7 (Brown , 1956 ) 0.7 8 6 MI D leptus 0.64 - 26-3 4 4.5-5. 7 11-1 2 51-5 6 OF F 8- 9 LS C 23-2 7 PO S 50-5 8 CO N BLP-AN N 3.0-4. 0 p p (AUen, 1955 ) 0.9 6 6 lineatus 0.85 - 30-3 3 4.8-5. 0 12-1 3 51-5 3 OF F 6 IN C 25-2 8 PO S 42-44 SC L HEM-SM O 3. 0 29 11 (Allen , 1955 ) 0.8 6 6 macrodens 1.02 - 30-3 2 5.1-6. 0 14-1 5 50-5 5 OF F 7 HS C 43-4 8 PO S 34-3 7 SCL SHM-SM O 2.6 37 11 (Allen , 1955 ) 1.12 n 6 macrurus 0.83 - 19-2 5 4.8-6. 3 14-2 0 54-5 9 CN T 8-10 ? HS C 25-3 4 PO S 6 39-4 7 CYL HEM-AN N 2.4 39 1 1 ( Goodey , 193 2 ) 1.19 microdorus 0.58 - 24-2 9 4.8-5. 9 11-1 3 55-5 9 OF F 4- 6 LS C 13-1 5 PO S 6 53-5 4 SCL BLP-SM O 2.5-3. 0 20-2 2 7- 8 (Geraert , 1966 ) 0.7 0 nanus 0.52 - 27-3 1 4.5-5. 3 10-1 2 52-5 7 CN T 7 LSC 12-1 5 PO S 6 55-6 0 CO N BLP-AN N 3. 8 23 8 (Allen , 1955 ) 0.6 4 noihus 0.55 - 24-3 0 4.0-5. 1 10-1 1 53-5 7 CN T 6 LS C 16-1 8 PO S 6 39-4 8 CO N SHM-AN N 3. 0 p p (Allen , 1955 ) 0.7 0 p p obscurisulcatus 0.7 9 50 5. 8 1 7 5 5 OF F 6 LS C 22 PO S 6 27-3 2 CO N BLP-SM O 3. 8 (Andrassy , 1959 ) obsciirus 0.63 - 26-3 7 4.6-5. 3 13-1 6 53-5 8 CN T 7 LS C 24-2 7 PO S 6 53-5 8 CO N BLP-AN N 3. 2 29 8

(Allen , 1955 ) 0.7 9 O3

F O l WASHI 140 PROCEEDINGS OF THE HELMINTHOLOGICAL SOCIETY

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Table 2. Departures from Tylenchorhynchus.

Tylenchorhynchus To By alattis (Cobb, 1930) species inquirendae Tarjan (1964) browni ( Kreis, 1929) species inquirendae Allen (1955) bucharicus (Tulaganov, 1949) species inquirendae Tarjan (1964) bnrsifer Loof, 1960 Paratrophurus Siddiqi (1971) caromatus (Tulaganov, 1949) species inquirendae Tarjan (1964) clavicaudatus Seinhorst, 1963 became clavicauda Seinhorst (1968) coffeae Siddiqi and Basir, 1959 species inquirendae Tarjan (1964) crassicaudatus Williams, 1960 syn. of mashhoodi Baqri and Jairajpuri (1970) dactylurus Das, 1960 syn. of mashhoodi Baqri and Jairajpuri (1970) digitatus Das, 1960 syn. of mashhoodi Baqri and Jairajpuri (1970) dissitus Colhran, 1969 Paratrophurus Siddiqi (1971) elegans Siddiqi, 1961 syn. of mashhoodi Baqri and Jairajpuri (1970) graminicolus Kirjanova, 1951 species inquirendae Tarjan (1964) indicus Siddiqi, 1961 syn. of brevilineatus Siddiqi (1963) mamillatus Tobar-Jimenez, 1966 Scutylenchus Jairajpuri (1971) ornatus Allen, 1955 syn. of quadrifer Andrassy in Tarjan (1964) paucus Kirjanova, 1951 species inquirendae Meyl (1961) rhopalocercus Seinhorst, 1963 Trichotylenchus Seinhorst (1968) sexamammillattts (Kirjanova, 1938) species inquirendae Tarjan (1964) spinicaudatus Sch. Stekhoven, 1944 Hirschmanniella Luc and Goodey ( 1963 ) styriacus ( Micoletzky, 1922 ) species inquirendae Allen (1955) symmetricus (Cobb, 1914) species inquirendae Allen ( 1955 ) trilincatus Timm, 1963 syn. of triglyphus Andrassy in Tarjan (1964) zeae Sethi and Svvarup, 1968 syn. of mashhoodi Baqri and Jairajpuri (1970)

Literature Cited species. (In Russian.) Tr. Karakalpaksk. Cos. Ped. In-T. 2: 175-185. Allen, M. W. 1955. A review of the nematode genus Tylenchorhynchus. Univ. Calif. Publ. Filipjev, I. N. 1936. On the classification of Zool. 61: 129-166. the Tylenchinae. Proc. Helm. Soc. Wash. 3: Andrassy, I. 1961. Wissenschaftliche Ergebnisse 80-82. der ersten ungarischen zoologischen Expedi- Geraert, E. 1966. On some Tylenchidae and tion in Ostafrika. 2. Nematoda. Ann. Hist.- Neotylenchidae from Belgium with the de- Nat. Mus. Nat. Hungar. n.s. 53: 281-297. scription of a new species, Tylenchorhynchus Arias-Delgado, Maria, F. Jimenez-Millan, and microdorus. Nematologica 12: 409-416. J. M. Lopez-Pedregal. 1965. Three new Goffart, H. 1930. Die Aphelenchen der Kul- species of possible phytoparasitic nematodes turpflanzen. Berlin, 105 p. in Spanish soils. (In Spanish.) Publ. Inst. Golden, A. M. 1971. Classification of the gen- Biol. Aplic. 38: 47-58. era and higher categories of the order Tylen- Baker, A. D. 1962. Check lists of the nema- chida (Nematoda), Chapt. 8, p. 191-232. In tode superfamilies Dorylaimoidea, Rhabditoi- B. M. Zuckerman et al. (eds.), Vol. I, Plant dea, Tylenehoidea and Aphelenchoidea. E. J. Parasitic Nematodes, Academic Press, New Brill, Leiden, Netherlands, 261 p. York. Baqri, Q. H., and M. S. Jairajpuri. 1970. On Goodey, T. 1932. The genus Anguillulina Gerv. the intra-specific variations of Tylenchorhyn- & v. Ben., 1859, vel Tylenchus Bastian, 1865. chus mashhoodi Siddiqi & Basir, 1959 and an J. Helminthol. 10: 75-180. emended key to species of Tijlenchorhynchus de Guiran, G. 1967. Description of two new Cobb, 1913 (Nematoda). Rev. Brasil. Biol. species of the genus Tylenchorhynchus Cobb, 30: 61-68. 1913 (Nematoda: Tylenchinae) with a key Braun, A. L., and P. A. A. Loof. 1966. Praty- to the females, and precisions on T. mamil- lenchoides laticauda n. sp., a new endopara- latus Tobar-Jimenez, 1966. (In French.) sitic phytonematode. Neth. J. Pi. Path. 72: Nematologica 13: 217-230. 2,41-245. Hopper, B. E. 1959. Three new species of the Eliashvili, T. S. 1971. Two new soil-inhabiting genus Tylenchorhynchus (Nematoda: Tylen- nematode species (Amphidelus paramonovi chida). "Nematologica 4: 23-30. and Tylenchorhynchus georgiensis n. sp.) of Izatullaeva, R. I. 1967. New nematode species eastern Georgia. (In Russian.) Bui. Acad. from ornamental flowering plants in Kazakh- Sci. Georgian SSR 61: 213-216. stan. (In Russian.) Izv. Akad. Nauk Kazakh. Erzhanova, P. K. 1964. Nine new nematode SSR, Ser. biol. Nauk 5: 45-50.

Jairajpuri, M. S. 1966. A redefinition of Psi- toda: Tylenchoidea). (In Russian.) Parazito- lenchus deMan, 1921 and Tylenchus sub- logiya 4: 342-344. genus Filenchus Andrassy, 1954 with the Netscher, C., and G. Germani. 1969. Telotij- erection of Clavilenchus n. subgenus under lenchus baoulensis n. sp. and Trichotylenchus Tijlenchus Bastian, 1865. Nematologica 11 rectangularis n. sp. (Nematoda, Tylenchoi- (1965): 619-622. dea). (In French.) Nematologica 15: 347- . 1969. On the identity of Telotylenchtis 352. Siddiqi, 1960 and Trichotylenchus White- Sabova, N. 1967. Two new soil inhabiting head, 1959 with remarks on their systematic nematode species (Tylenchorhynchus tatren- position. (Abst.) All India Nemat. Symp., sis and Alaimus andrassyi n. spp.) from IARI, New Delhi: 25. Czechoslovakia. Opusc. Zool., Bpest. 7: 237- . 1971. On Scutylenchus mamillatus (To- 240. bar-Jimenez, 1966) n. comb. (Abst.) Nat. Seinhorst, J. W. 1963. Five new Tylencho- Acad. Sci., India, 40th Session, p. 18. rhynchus species from West Africa. Nema- 1971. On the synonymy of Telotylen- tologica 9: 173-180. chus Siddiqi, 1960 with Trichotylenchus . 1968. Trichotijlenchus rhopalocercus Whitehead, 1959 (Nematoda: Tylenchida). (Seinhorst, 1963) n. comb. (syn. Tylencho- Indian J. Nematol. 1: 3-6. rhynchus rhopalocercus Seinhorst, 1963) and Kirjanova, E. S. 1951. Soil nematodes found Tylenchorhijnchus clavicauda nom. nov. (syn. in cotton fields and in virgin soil of Golod- T. clavicaudatus of Seinhorst, 1963). Nema- naya Steppe ( Uzbekistan ) . (In Russian. ) tologica 14: 596. Trudy Zool. Inst. Akad. Nauk SSSR 9: 625- 1971. On the genera Trichotijlenchus 657. and Telotijlenchus. Nematologica 17: 413— Knobloch, Natalie A. 1971. Emendation of 416. the description of Tylenchorhynchus longus Wu, 1969. Nematologica 17: 602-603. Seshadri, A. R., T. S. Muthukrishnan, and S. Litvinova, N. F. 1946. Four new species of Shunmugam. 1967. A new species of Tylenchorhynchus ( Nematoda ) from Kazakh- Tt/lenchorhynchus (Tylenchidae: Nematoda) stan. Proc. Zool. Soc. London 116: 120-128. from Madras State, India. Curr. Sci. 36: Loof, P. A. A. 1960. Miscellaneous notes on 551-553. the genus Tylenchorhynchus ( Tylenchinae : Sethi, C. L., and G. Swarup. 1968. Plant para- Nematoda). Nematologica 4 (1959): 294-306. sitic nematodes of northwestern India. I. The Luc, M., and J. B. Goodey. 1963. Hirschman- genus Tylenchorhynchus. Nematologica 14: niella nom. nov. for Hirschmannia. Nemato- 77-88. logica 9: 471. Sher, S. A. 1968. Revision of the genus Radoph- Merny, G. 1964. A new Tylenchida from trop- olus Thorne, 1949. (Nematoda: Tylenchoi- ical Africa: Tetylenchus annulatus n. sp. (In dea). Proc. Helm. Soc. Wash. 35: 219-237. French.) Nematologica 10: 425-430. . 1970. Reclassification of the genus Chi- - , and G. Germani. 1968. Tylenchorhyn- tinotylenchus (Micoletzky, 1922) and a re- palustris n. sp. (Nematoda: Tylenchi- description of C. paragracilis (Micoletzky, nae) inhabitant of the rice fields of the Ivory 1922) (Nematoda: Tylenchoidea). J. Nem- Coast. (In French.) Ann. Epiphyties. 19: atol. 2: 236-238. 601-603. Siddiqi, M. R. 1963. Four new species in the Meyl, A. W. 1961. Die freilebenden Erd- und sub-family Tylenchinae (Nematoda) from Svisswassernematoden (Fadenwiirmer). In Die North India. Z. Parasitenk. 23: 397-404. Tierwelt Mitteleuropas, Brohmer P., P. Ehr- . 1970. On the plant-parasitic nematode mann, and G. Ulmer ( ed. ) . Quelle & Meyer, genera Merlinius gen. n. and Tylenchorhijn- Leipzig, 273 p. chus Cobb and the classification of the fami- Micoletzky, H. 1922. Die freilebenden Erd-Ne- lies Dolichodoridae and Belonolaimidae n. matoden — und Bestimmungsschliisseln. Arch. rank. Proc. Helm. Soc. Wash. 37: 68-77. Naturg., Berlin (1921), Abt. A, 87: 321- 1971. Structure of the oesophagus in 650. the classification of the superfamily Tylen- Mulvey, R. H. 1969. Nematodes of the genus choidea (Nematoda). Indian J. Nematol. 1: Tylenchorhynchus (Tylenchoidea : Nematoda) 25-43. from the Canadian high Arctic. Can. J. Zool. Steiner, G. 1914. Freilebende Nematoden aus 47: 1245-1248. der Schweiz. 1. Teil einer vorlaufigen Mit- Mukhina, T. I. 1970. A new species of the teilung. Arch. Hydrobiol. 9: 259-276. genus Ttjlenchorhynchus Cobb, 1913 (Nema- . 1920. Freilebende Siisswassernematoden

aus peruanischen Hochgebirgsseen. Revue Tobar-Jimenez, A. 1970. Description of two Suisse Zool. 28: 11-44. new species of the genus Tylenchorhynchus Sturhan, D. 1966. Ueber Verbreitung, Patho- Cobb, 1913 (Nematoda: Tylenchidae) with genitat und Taxonomie der Nematodengat- some additional data on T. sulcatus de Gui- tung Tylenchorhynchus. Mitt. Biol. Bund. ran, 1967. (In Spanish.) Rev. Iber. Para- Land- u Forst. Berlin-Dahlem. 118: 82-99. sitol. 30: 215-228. Szczygiel, A. 1970. Tylenchorhynchus polonicus Wallace, H. R., and D. N. Greet. 1964. Ob- sp. n. and Helicotylenchus pseudodigonicus servations on the taxonomy and biology of sp. n. (Nematoda, Tylenchoidea) from Po- Tylenchorhynchus macrurus (Goodey, 1932) land. Bull. Acad. Pol. Sci. Cl. V 17: 685-690. Filipjev, 1936 and Tylenchorhynchus icarus Tarjan, A. C. 1964. A compendium of the sp. nov. Parasitology 54: 129-144. genus Tylenchorhynchus (Tylenchidae: Ne- Whitehead, A. G. 1959. Trichotylenchus fal- matoda). Proc. Helm. Soc. Wash. 31: 270- 280. ciformis n. g., n. sp. (Belonolaiminae n. sub- Thome, G. 1949. On the classification of the farn.: Tylenchida Thorne, 1949) an associate Tylenchida, new order (Nematoda, Phasmi- of grass roots (Hypanhenia sp.) in southern dia). Proc. Helm. Soc. Wash. 16: 37-73. Tanganyika. Nematologica 4: 279-285. , and R. B. Malek. 1968. Nematodes Wu, L. Y. 1969. Three new species of the of the northern great plains. Part 1. Tylen- genus Tylenchorhynchus Cobb, 1913 (Tylen- chida (Nemata: Secernentea). So. Dakota chidae: Nematoda) from Canada. Can. J. Agr. Exp. Sta. Tech. Bull. 31, 111 p. Zool. 47: 563-567.

Plagioporus hypentelii sp. n. (Trematoda: Opecoelidae) from the Hogsucker, Hypentelium nigricans (LeSueur) (Osteichthys: Catostomidae)1

SHERMAN S. HENDRix2 Department of Zoology, University of Maryland, College Park, Maryand

ABSTRACT: Plagioporus hypentelii sp. n. (Trematoda: Opecoelidae) is described from the intestine of the hogsucker, Hypentelium nigricans (LeSueur), in streams of Adams County, Pennsylvania. It most closely resembles P. serotinus Stafford, 1904, but differs from it chiefly in its less elongate shape, smaller size, smaller testes, ovary, and eggs, straight cirrus sac, in hosts and geographic location of the hosts.

During the course of examination of fresh- few hours of capture. The worms were studied water fishes for endohelminths, an unde- alive and then fixed in hot AFA with slight scribed trematode of the genus Plagioporus coverslip pressure. Whole mounts were stained Stafford, 1904, was recovered from the intes- with Semichon's acetocarmine and mounted tine of the northern hogsucker, Hypentelium in permount. Sections were stained with nigricans (LeS ueur). Harris' hematoxylin and eosin. Fish hosts were collected with nets or by Unless stated otherwise, the following de- electrofishing and usually autopsied within a scription is based upon 25 stained whole mounts and five sectioned specimens. All 1 From a dissertation submitted to the Graduate School, measurements are in micrometers. Means are University of Maryland, in partial fulfillment of the require- ments for the Ph.D. degree in Zoology. given first followed by the range in parentheses 2 Present address: Department of Biology, Gettysburg College, Gettysburg, Pennsylvania 17325. and are from the whole mounts.